Herpetology Notes, volume 14: 1027-1035 (2021) (published online on 03 August 2021)

On the second (or, rather, the first) specimen of the recently described salitan (: ), with the first report of the from Island, southern

Rafe M. Brown1,*, Marta Calvo-Revuelta2, Johana Goyes Vallejos3, Alberto Sánchez-Vialas2, and Ignacio De la Riva2

Abstract. We report on the second known specimen of the recently described banded Asian coralsnake, Calliophis salitan, identified among the Museo Nacional de Ciencias Naturales collections of Dr. Hipólito Fernández, made in1886.The specimen was collected on Mindanao Island in the southern Philippines as part of the “Comisión Central de Manila,” the Spanish Crown’s effort to catalogue and showcase Philippine biodiversity for the Madrid public. We assign this important specimen to C. salitan (formerly known from a single specimen collected on ) based on its highly distinctive external phenotypic characteristics (scalation, body size, and inferred colour pattern). The discovery of C. salitan on Mindanao significantly extends the known geographic range of this species and partially fills an intermediate distributional hiatus – a large geographic gap in the distribution of large-bodied, long-glanded coralsnakes. This report partially resolves a biogeographic enigma surrounding the question of how a distinctive evolutionary lineage of elapid – more closely related to the Calliophis bivirgatus group from Borneo and other landmasses of Sundaland than to any other Philippine elapids – could have colonized only a small island (Dinagat) in the geographic interior of the archipelago. The answer appears, most likely, that this dispersal occurred over land, via the large intervening island of Mindanao. Although the exact site of collection is unknown, consideration of the collector’s itinerary suggests that it originated most likely in the vicinity of Davao City, southeastern Mindanao. Its discovery confirms that C. salitan occurs on Mindanao (or, at least, did occur on this island as recently as 133 years ago), and that locating additional populations via targeted surveys is an urgent priority for conservation of this remarkable Philippine endemic lineage.

Keywords. Coralsnakes, distribution, Elapidae, , Southeast Asia.

Introduction Conceptually, these topics invoke a number of evolutionary questions, concerning phenomena not well Compared with neighbouring Sundaland, the understood. Philippines has a relatively depauperate terrestrial elapid The first of these is the unexplained and evolutionarily snake fauna. Terrestrial Philippine snakes of the family anachronistic case of sexual size dimorphism in the Elapidae (Leviton, 1964; Leviton et al., 2014, 2018) diameter of the discharge orifice in venom-conducting include Ophiophagus (one species), (three species), fangs of Naja philippinensis, which is larger in females Hemibungarus (three species), and Calliophis (four (Wüster and Thorpe, 1992). This is the only known case species). Despite the relatively low diversity of land- of this phenomenon in cobras, and such a disparity may dwelling elapid species in the archipelago, the Philippine be expected to have functional significance (Bogert, elapid snake fauna possesses interesting attributes in 1943). However, an unambiguous explanation of its terms of natural history, ecology, and biogeography. adaptive significance in Philippine cobras has yet to be provided (Wüster and Thorpe, 1992). Second, on a related point, Philippine cobras possess 1 Biodiversity Institute and Department of Ecology and highly-specialized spitter dentition (rifled, mobile fangs Evolutionary Biology, University of Kansas, Lawrence, for frontal projection of venom; Wüster and Thorpe, Kansas 66045, USA. 2 Museo Nacional de Ciencias Naturales (MNCN-CSIC), Calle 1992), and yet conflicting accounts and ambiguous José Gutiérrez Abascal 2, 28006, Madrid, Spain. reports of actual spitting behaviour in Philippine 3 Division of Biological Sciences, University of Missouri, “spitting” cobras N. philippinensis and N. samarensis Columbia, Missouri 65211, USA. (Taylor, 1922, 1975; Leviton, 1964, Leviton et al., 2014) * Corresponding author. E-mail: [email protected] suggest an unresolved disconnect between structure and © 2021 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0. function (Watt et al., 1988; Wüster and Thorpe, 1991, 1028 Rafe M. Brown et al.

1992). In addition to unclear literature accounts, we Aggregate Island Complexes, or PAICs – sensu Brown have never observed spitting behaviour in these species, and Diesmos, 2009; Brown et al., 2013a), whereas four despite numerous field encounters over the past three PAIC-endemic species of Calliophis are restricted to decades (RMB, pers. obs.). three faunal regions of the southern and western reaches Third, Philippine elapids possess unexpected and of the archipelago (Fig. 1; Leviton et al., 2018). surprising phylogenetic relationships. The close, Finally, snake biogeographers have been puzzled by sister-lineage relationship between king cobras of the the absence of kraits and large-bodied, brightly-coloured Ophiophagus and the Philippine endemic false coralsnakes in the archipelago. These conspicuous coralsnakes of the genus Hemibungarus was entirely faunal elements are well-documented and frequently unexpected because of their remarkably dissimilar encountered just outside of the Philippines (on the phenotypes, anatomy, ecology, microhabitats, and islands of Borneo, Sumatra, Java, Sulawesi, and the behaviour (Castoe et al., 2007; Brown et al., 2018; Thai-Malaysian Peninsula of mainland Southeast Asia; Plettenberg Laing, 2018). Uetz et al., 2020), but have been notably absent within Fourth, the biogeography of Philippine elapids is the archipelago (but see below). interesting because it involves a peculiar geographic The first four topics summarized above represent replacement pattern of coralsnakes (Leviton, 1964; reasonably well understood or, at least, partially Brown et al., 2018). In this case, the three endemic documented Philippine-specific elapid case histories that species of Hemibungarus have come to occupy three can be viewed as intriguing challenges for international separate faunal regions of the northern, northeastern, collaborations involving Philippine scientists, students, and central portions of the archipelago (the Pleistocene and researchers from nearby Southeast Asian countries

Figure 1. Map of the southern Philippines with the specimen-vouchered distributions of Philippine Calliophis species indicated by various symbols as shown in the key. A yellow star marks the C. salitan type locality on northern Dinagat Island. According to the itinerary of Hipólito Fernández, collector of the first specimen ofCalliophis salitan from Mindanao Island (MNCN 22719), he accessed the island through the Port City of Davao (orange star). First Report of Calliophis salitan from Mindanao Island 1029 and beyond (Alcala, 2004a, b). However, the last point questions (Brown et al., 2018). One of these we address enumerated above has recently been addressed with the here, with the first report of a historically-significant, discovery of a strikingly distinct, spectacularly-coloured, unrecognized specimen, which the available evidence new species of long-glanded coralsnake, Calliophis suggests likely originated on the intervening island of salitan Brown et al., 2018, heretofore known from a Mindanao. The discovery of the second specimen of C. single specimen. This species, documented only from salitan, reidentified 133 years after the first record of its Dinagat Island northeast of Mindanao (Fig. 1), remains existence, and housed within the historic herpetological enigmatic because virtually nothing is known about its collections of Spain’s National Museum, the Museo biology. Calliophis salitan was discovered in 2007 and Nacional de Ciencias Naturales (MNCN), constitutes a was described on the basis of a single specimen (Brown et significant biogeographical “piece of the puzzle” in the al., 2018). Related species are referred to as “long-glanded sense that it fills a distributional gap between the central tropical coralsnakes” because they possess remarkably Philippine island of Dinagat, where C. salitan was enlarged venom glands, which extend from the head to discovered, and the Sunda Shelf landmasses of Borneo, the thoracic-cardiac cavity (Brown et al., 2018: Fig. 6). Java, Sumatra, and the Thai-Malaysian Peninsula, In the years since the initial discovery of C. salitan where its closest relatives are found (Stuebing et al., (by J.B. Fernandez in 2007) and its description (Brown 2014; Charlton, 2020). et al., 2018; Leviton et al., 2018; Weinell et al., 2019), numerous biodiversity surveys and faunal inventories Historical background and Philippine herpetological of Dinagat Island (as well as nearby Island), collections at MNCN: The “Comisión Central de southern and islands, to the north, and Manila” of 1885. In his MNCN catalogue of Philippine northeastern Mindanao and Sur islands to vertebrate specimens collected prior to 1886 and the south (Fig. 1) have been conducted by qualified assembled in 1887 as part of the Comisión’s activities, herpetologists (see Sanguila et al., 2016 for a review). Gogorza (1888) sought to summarize all that was known of However, the search for additional specimens of C. specimen-based vertebrate biodiversity of the Philippines, salitan, or clues to the extent of its geographical range, for later presentation to the scientific and lay public of its biogeography, and natural history (diet, ecology, Spain. This catalogue summarized each specimen’s microhabitat preference, venom toxicity, etc.) have identity, the locality from which it had been secured, the all been fruitless. Eventually, after considerable but collector, and the date. The Comisión was atypical in the unsuccessful efforts to secure additional specimens, sense that no standard expeditions were carried out during the species was described (Brown et al., 2018) and a or leading up to 1887. Instead, an agreement was made molecular phylogenetic analysis demonstrated its close to first hold an exhibition covering aspects of history relationships to the large-bodied (≥ 1 m total body (including natural history) of the Philippines in-country, length), active “racer ecomorph,” long-glanded and and then subsequently transport this exhibition to Madrid, spectacularly-coloured (blue-black iridescent bodies, in a manner similar to public exhibitions that were the bright orange-red heads and tails) coralsnakes of trend, hosted by other European countries and colonies at Sundaland: the Calliophis bivirgatus complex, which the time. To do this, a “Comisaría Regia” was constituted includes the taxa C. b. bivirgatus, C. b. flaviceps, and in Madrid, and given the responsibility of organizing C. b. tetrataenius. In terms of their size, morphology, both the exhibition site in Madrid and also the Comisión colouration, ecology, behaviour, and temperament, the (located in the Philippine capital city of Manila on the Sundaland large-bodied, coralsnakes of the C. bivirgatus island of ). The latter was charged with gathering complex are much more reminiscent of kraits (e.g., all possible zoological specimens, collected throughout Bungarus flaviceps; see Stuebing et al., 2014) than they the archipelago by various collectors spanning different are of the contrastingly secretive, small-bodied, rarely traditions and trades (physicians, engineers, naturalists, observed, inconspicuously coloured, and semi-fossorial monks, priests, etc.) that would subsequently be brought Philippine species of the C. intestinalis complex, which to Madrid (Catálogo, 1887), thereby assembling a includes C. bilineatus from the PAIC, C. collection of hundreds of specimens. philippinus from the Mindanao PAIC, and C. suluensis The main contributors to the Comisión’s collecting from the Sulu Archipelago PAIC. activities were Hipólito Fernández and Domingo Sánchez The surprising discovery of a species of the C. bivirgatus y Sánchez. Other contributors included José Pérez Maeso, complex on Dinagat Island left several unanswered Carlos Mazarredo, Ramón Jordana, Agustín Domec, 1030 Rafe M. Brown et al.

Regino García, and Claudio Montero. A summary of the Antipolo (Morong); Daraga, Irosin, and Tabaco cities numbers and locations of their contributions follows. (vicinity of Mayon Volcano, Albay Province); Negros The prominent physician and collector Hipólito Island, vicinity of Dumaguete City (West Visayas); Leyte Fernández collected on Luzon near Manila, and also Island (East Visayas); Island, vicinity of Boac near the port at Lagonoy City (Camarines Sur Province), City; Island. Ramón Jordana y Morera, also Lobo (Batangas Province), Mindoro Island, and on some a forest engineer, only collected on (south of the West Visayan islands. He also visited Mindanao of Palawan). MNCN specimens include three specimens Island by way of the port city of Davao. Fernández of Lycodon muelleri and two Dendrelaphis marenae. collected a total of 130 reptiles (91 specimens of 33 Agustín Domec, Navy officer and physician, resident snake species, all in collections at MNCN), including in the Philippines, appointed by MNCN’s “Facultative MNCN 22719, a large elapid snake reportedly from “Isla Board” as collaborator, collected six specimens of snakes de Mindanao, Filipinas,” (date of specimen cataloguing on Mindanao, presumably in the vicinity of Davao City 30 June 1887; presumably collected sometime before (1881), representing five species (notably including a this date) but with no more specific locality information, specimen of Naja samarensis; reported as N. tripudians which is the subject of this report. by Gogorza, 1888: 279), all of which are Mindanao faunal Domingo Sánchez y Sánchez, for whom an 1886 region endemics or widespread species found throughout itinerary is available in the MNCN archives, embarked the archipelago (Leviton et al., 2018). Claudio Montero, aboard a paddle steamer, the Queen Isabel II, from the Navy Rear Admiral, collected in the Philippines (but port of Manila on 10 January 1886 and travelled to without detailed locality information), and contributed Bulacán, returning on 10 February. Localities visited 58 reptiles, including 34 snake specimens, representing included Quiñgua, Montes Angat (vicinity of today’s 16 species. Angat Dam), Pinubayan Forest (vicinity of the Constancia In summary, a total of 336 snakes, comprising 49 species, Mine) and Tayabas. Between February and March 1886, were deposited in the MNCN Philippine collection by he conducted an expedition to Samar Island, directed by 1885 (González-Fernández, 2002). Subsequently, in D. Sebastián Vidal, together with ichthyologist Regino 1887, this collection was first celebrated as part of the García. This excursion collected in Paranas, Villareal, “Exposición General de las Islas Filipinas”, exhibited in Zumarraga, Daram, Bioso, and Catbalogan. In total 77 the Parque El Retiro. On the occasion of this exhibition, specimens of reptiles were collected (38 snakes of 15 a new palace, the “Palacio de Cristal” (formerly known species) from Samar localities and the following sites as “pabellón estufa”) was constructed, and served as a on Luzon: Manila, Lobo and Nasugbu, Tuy, and Lian greenhouse public exhibition, featuring live Philippine (Batangas Province), Dolores (Tayabas), Angat and collected flora (no MNCN zoological specimens were Quiñgua (Bulacan), Morong and Bosoboso (Rizal), and exhibited). Once the exhibition was finalized in 1887, the Calauan and Los Baños (Laguna). nearby Museo de Ultramar (located in the current Palacio José Pérez Maeso collected Philippine specimens de Velázquez), housed the Philippine MNCN zoological before the Comisión was established. In 1885 he made a collections (Gogorza, 1888), until it was decommissioned donation to the MNCN (Barreiro, 1992) of 31 specimens in 1908, and Philippine specimens were transferred back of reptiles, including 14 snakes (10 species). He collected to MNCN. A total of 39 elapid snakes, representing 12 on Mindanao (in Surigao Province and on the Zambonaga species from Philippines, are conserved at MNCN; these Peninsula) and other islands, including southern include the marine elapid, seasnake species Hydrophis Island, Samar, “Isla de Paragua” (= Palawan Island, schistosa, H. inornatus, H. fasciatus, H. hardwickii, and vicinity of Puerto Princesa City), and on Luzon (in Pelamis platurus, the freshwater H. semperi, specimens Bulacan Province). of the terrestrial elapid cobra species Naja samarensis, Carlos Mazarredo, forest engineer, donated a collection and exemplars of the terrestrial coralsnake species of a total of 76 specimens of reptiles, including 34 snake Hemibungarus calligaster, H. mcclungi, H. gemianulis, specimens representing 15 species from various islands and Calliophis philippinus. Additionally, a single (all collected by him during 1884–1885; Barreiro, 1992: noteworthy specimen of Calliophis was housed in the 347). These included specimens from Luzon: Dolores collection. We examine the morphology of this specimen (Tayabas Province); vicinity of Bulacan, Cuevas de Puning and discuss its taxonomic identity and relevant historic (Bulacan Province); vicinity of Bula and Bato Cities, implications below. and from Nueva Cáceres (Camarines Sur Province); First Report of Calliophis salitan from Mindanao Island 1031

Materials and Methods possesses 13 rows of dorsals that remain unreduced along the entire body (vs. 17–25 in Naja, 15 in Ophiophagus and We examined all preserved specimens of the genus Hemibungarus), and has a single primary temporal scale Calliophis from the Philippines found in the MNCN. (vs. two in species of Hemibungarus; Weinell et al., 2019: We obtained measurements of external morphology Fig. 17C). Thus, MNCN 22719 cannot be confused with directly from the specimens and follow the standard any other species of elapid snake from the Philippines and terminology and methods for quantifying and is unambiguously identified here asC. salitan. describing of elapid snake scalation of Brown et al. MNCN 22719 has a total length of 998 mm, snout– (2018). We followed Weinell et al. (2019) to identify vent length of 841 mm, and tail length of 157 mm. The key characteristics distinguishing species of the genera specimen’s hemipenes were not everted at preservation Calliophis, Naja, and Hemibungarus, and once we but sex was confirmed by probing; slight lateral bulges, unambiguously ruled out the possibility that MNCN interpreted as hemipenial swellings, are evident in the 22719 could be a member of any other elapid genus, ventral aspect at the tail base (Figs. 2C, D). All dorsal we used the diagnosis of Brown et al. (2018) and the body scales are unkeeled, in 13 rows, and unreduced illustrated key provided by Weinell et al. (2019) to for the entire length of the body. MNCN 22719 has 257 confirm species identity. Measurements were taken vertebral scales, counted from the parietals to a point with a flexible fabric measuring tape (by RMB, to the on the dorsal tail base at the level corresponding to the nearest mm), and scale characters were scored with posterior edge of the precloacal shield, and then 60 dorsal the naked eye. A complete list of all other Philippine vertebral caudal scales to the end of the tail (not including elapid specimens examined was provided by Brown et the tail tip), and 58 subcaudals. MNCN 22719 has 6/6 al. (2018) and we consulted that work, as well as that of (left/right) supralabials, and 7/7 infralabials. Prefrontals Leviton et al. (2018), for recent summaries of diversity are slightly wider than long, in contact laterally with the and documented distributions of all other Philippine nasal, preocular, and supraocular scales (Fig. 3B); the snakes of the family Elapidae. specimen has 1+0 temporal and a single post-temporal, a single preocular, two postoculars (the lower is less Results than half the size of the upper), and no loreal scales (i.e., Among the Museo Nacional de Ciencias Naturales the nasal and the preocular are in contact; Fig. 3A). The collections of Hipólito Fernández is a single specimen postnasal is triangular and the first pair of infralabials of Calliophis salitan, accompanied by data indicating is in contact behind the mental, whereas the second “30 June 1887 from Isla de Mindanao, Filipinas, by H. pair is small and separated. The second and third pair Fernández (No. Col: 7424[P]).” The specimen, an adult of infralabials touch the anterior-most elongate pair of male, is intact, in reasonably good condition, with four genials (Fig. 3C); the fourth pair are largest and contact 20–30 mm incisions on the ventrum (through ventrals the anterior and posterior genials and the first sublabial. but not subcaudals). MNCN 22719 (Figs. 2, 3) was The fifth–seventh infralabials contact the first sublabial, presumably collected some time before the date of its which is in contact with the genials. Three scales separate accessioning, and it was initially identified by Gogorza the posterior genials and the first ventral (Fig. 3C). The (1888) as Naja sp., then later reidentified and reported as first of these is a small, non-expanded gular scale. The N. naja samarensis (González-Fernández, 2002). At some second and third are increasingly laterally expanded point over the last two decades, its identification was gulars (or “preventrals”). Subcaudals are divided but the changed in the MNCN database to . precloacal scale is single (Fig. 2D). However, even without pigmentation characters, the high number of subcaudals, and the relatively long tail (relative Discussion to total body length), rules out any possibility that MNCN Comparison of MNCN 22719 to other Mindanao 22719 could be conspecific with any of theC. intestinalis populations of small-bodied, long-glanded coralsnakes group species (Leviton, 1964; Brown et al., 2018). reinforced our conclusion that the specimen cannot be Additionally, MNCN 22719 possesses two postoculars confused with any other Philippine species. Mindanao (whereas three are present in Naja and Ophiophagus; specimens considered conspecific with Calliophis Weinell et al., 2019: Figs. 17A, B), has a distinctly intestinalis until the 1850s were originally described triangular postnasal (Fig. 3A) vs. the vertically elongate as “Callophis intestinalis var. Philipp.” by Günther postnasal typical in Naja (Weinell et al., 2019: Fig 17A), (1859: Plate XVI, Fig. A). However, despite the detailed 1032 Rafe M. Brown et al.

Figure 2. Two specimens of Calliophis salitan from the southern Philippines. (A) The holotype from Dinagat Island in dorsal view. (B) The holotype in ventral view. In life, the tail was bright orange but has faded since preservation in 2007. (C) The Mindanao specimen (MNCN 22719) in dorsal view. (D) The same specimen in ventral view. Note the faint pigmentation, suggestive of the species’ diagnostic colour pattern: broad, alternating black and white body bands. Photos by Rafe Brown (A, B) and Alberto Sánchez-Vialas (C, D). description this earliest name for a distinct Philippine Based on his itinerary, we interpret Fernández’s population has no nomenclatural validity because of the Mindanao specimen (MNCN 22719) as likely having abbreviated, non-Latinised third part of the name. It was been acquired in the vicinity of Davao City, in the only validated as Callophis intestinalis var. philippina southeastern portions of the island (Fig. 1). Although by Günther (1864: 349). Subsequently, it was treated by the hemipenes of MNCN 22719 were not everted at Leviton (1964) as the subspecies Maticora intestinalis preservation (see Brown et al., 2018: Fig. 7C), we non- philippina and later transferred back to Calliophis invasively confirmed sex, and additionally emphasize following phylogenetic analyses of anatomical characters that high subcaudal counts in MNCN 22719 (even higher (Slowinski et al., 2001). They were subsequently elevated than the demonstrably male holotype – the only other to the level of full species, as C. philippinus (Leviton known specimen), support our determination of sex; et al., 2014, 2018; Sanguila et al., 2016; Brown et al., no females have yet been reported. The body size and 2018). Calliophis philippinus is a much smaller species proportions of the Mindanao specimen nearly match (the largest male recorded had a total length of 713 mm, those of the holotype (Brown et al., 2018). It is nearly the largest female 575 mm; Brown et al., 2018: Table 3), 1 m long (total length 998 mm), with a notably long tail and the majority of specimens available are typically that comprises 15.6% of the total length (14.1% in the less than half the body length and diameter of C. salitan holotype) and provides an important diagnostic character (RMB, pers. obs.). In contrast, C. salitan is a much larger that distinguishes C. salitan from all relevant congeners. , with a total body length ≥ 1 m. As inferred in a recent molecular phylogenetic analysis First Report of Calliophis salitan from Mindanao Island 1033

has 235; ventrals number 249 in the Mindanao specimen, and 229 in the holotype. The Mindanao specimen has 60 caudals and 58 subcaudals (the holotype has 55 caudals and 54 subcaudals). In all other standard meristic variables and categorical scalation characters, the two specimens are remarkably consistent. In the last 133 years, MNCN 22719 has faded considerably, with almost all presumed colour pattern features (bright red tail; neck and remainder of dorsal surfaces of body with alternating, broad, black and white bands; dorsal head black; Brown et al., 2018: Figs. 4A–C, 5A–B) faded, presumably as a result of exposure to ultraviolet light, either while on public exhibit during the years 1887–1889, or beyond. Nevertheless, faintly evident are at least eight faded but discernible, broad, dark transverse body bars (Fig. 2C), matching the documented colour pattern of this distinctive species, which cannot be confused with any other known Philippine elapid snake (Brown et al., 2018; Fig. 2). The holotype, PNM 9844, is also a male, with ten dorsal body bands including the nuchal band (which is continuous with dorsal black cephalic colouration), a creamy off-white throat and chin, but the remaining ventral surfaces of the body have nine broad transverse bands (corresponding to the remainder of those on the dorsum). Ventral surfaces of MNCN 22719 are immaculate (Fig. 2D), and it is unclear whether dorsal bands originally had corresponding ventral colour elements as in the holotype (Fig. 2A, B; Brown et al., 2018: Fig. 5B). Nevertheless, the width of both specimens’ transverse bands agrees sufficiently Figure 3. Lateral (A), dorsal (B), and ventral (C) views of the (corresponding to ~14–17 paravertebral scale rows). head of Calliophis salitan from Mindanao (MNCN 22719). In summary, the reidentification of MNCN 22719 as an Photos by Alberto Sánchez-Vialas. unmistakable second example of the enigmatic Philippine long-glanded coralsnake, Calliophis salitan, provides resolution for a biogeographic quandary (for discussion, (Brown et al., 2018), C. salitan is most closely related see Brown et al., 2018) by filling a puzzling distributional to the other large-bodied, non-Philippine long-glanded hiatus, with this new record from Mindanao Island. How coralsnakes, C. bivirgatus, and C. intestinalis. These, the species has gone unrecorded for over more than a and also the small-bodied, related congeners from the century of herpetological survey work in the Mindanao Philippines (C. bilineatus, C. philippinus, C. suluensis), faunal region (Taylor, 1922; Rabor and Alcala, 1959; all possess markedly shorter relative tail lengths (3.7– Leviton, 1963; Brown and Alcala, 1970; Ross and Lazell, 11.1% for both sexes of all three species). The Mindanao 1991; Sanguila et al., 2016; Brown et al., 2018) at first specimen’s snout–vent length is 841 mm, similar to that appears inexplicable. However, when one considers the of the holotype (PNM 9844 SVL = 856 mm) and its tail brief, unsustained, and for the most part ancillary nature length is 157 mm (tail length = 141). of most herpetological surveys over the last 133 years in Only a few notable differences in standard scalation the southern Philippines (Smith, 1993a,b; Delima et al., meristic characters are evident between the two male 2006; Nuñeza et al., 2010; Supsup et al., 2017), a likely specimens. The Mindanao specimen (MNCN 22719) explanation emerges. Rare, elusive, or secretive snake has 257 vertebrals, whereas the holotype (PNM 9844) species, such as arboreal or fossorial taxa, are difficult 1034 Rafe M. Brown et al. to detect and, when discovered during a survey, are Island, II: preliminary report on the herpetofauna of Aurora typically represented by single or very few specimens. Memorial National Park, Philippines. Hamadryad 25: 175–195. At naturally low abundances, such species may require Brown, R.M., Siler, C.D., Oliveros, C.H., Esselstyn, J.A., Diesmos, A.C., Hosner, P.A., et al. (2013a): Evolutionary processes of extensive and continuous surveys to be detected (Ferner diversification in a model island archipelago. Annual Review of et al., 2000; Gaulke, 2011; Sanguila et al., 2016), and Ecology, Evolution, and Systematics 44: 411–435. even then these species may only be represented by few Brown, R.M., Siler, C.D., Oliveros, C.H., Welton, L.J., Rock, A., specimens in collections or even singletons (Brown et al., Swab, J., et al. (2013b): The amphibians and reptiles of Luzon 2000; Siler et al., 2011; Brown et al., 2013b; Wynn et Island, Philippines, VIII: the herpetofauna of Cagayan and al., 2016; Weinell et al., 2020). It has become abundantly Isabela Provinces, northern Sierra Madre Mountain Range. clear that sustained or repeated, survey-and-resurvey Zookeys 266: 1–120. Brown, R.M., Smart, U., Leviton, A.E., Smith E.N. (2018): A new sampling efforts are necessary to comprehensively species of long-glanded coralsnake of the genus Calliophis characterize faunal diversity in the complex forests of the (Squamata: Elapidae) from Dinagat Island, with notes on the Philippines (Sison et al., 1995; Brown et al., 2000; Ferner biogeography and species diversity of Philippine Calliophis et al., 2000; Gaulke, 2011; Siler et al., 2011; Sanguila and Hemibungarus. Herpetologica 74: 89–104. et al., 2016). By inventorying species diversity across Brown, W.C., Alcala, A.C. (1970): The zoogeography of the elevational gradients, with attention to sampling across Philippine Islands, a fringing archipelago. Proceedings of the atmospheric and temporal/seasonal variation (e.g., dry California Academy of Sciences 38: 105–130. Catálogo (1887): Catálogo de la Exposición General de las Islas season versus wet season), sampling in different major Filipinas; Inaugurada por S. M. la Reina Regente, Celebrada en habitat types, and with particular attention to structural Madrid el 30 de Junio de 1887. Madrid, Spain Establecimiento microhabitat diversity, future field biologists may come Tipográfico de Ricardo Fé. to understand the natural history of the elusive C. salitan. Castoe, T.A., Smith, E.N., Brown, R.M., Parkinson, C.L. (2007): Higher-level phylogeny of Asian and American coralsnakes, Acknowledgements. We thank the U.S. National Science their placement within the Elapidae (Squamata: Serpentes), Foundation for financial support (DEB 0743491 and 1654388 and the systematic affinities of the enigmatic Asian coralsnake to RMB) that led to the discovery and description of C. salitan, Hemibungarus calligaster (Wiegmann, 1834). Zoological and U. Smart, A. Leviton, and E. Smith for collaboration on its Journal of the Linnean Society 151: 809–831. description. We also thank J. Fernandez, J. Weinell, C. Siler, Charlton, T. (2020): A Guide to the Snakes of Peninsular M.B. Sanguila, and A.C. Diesmos for their collaborative efforts and . Kota Kinabalu, Malaysia, Natural History in support of our work on Philippine elapids. RMB acknowledges Publications (Borneo). funding from the Research Excellence Initiative of KU’s College Delima, E.M.M., Ates, F.B., Ibañez, J.C. 2006. Species of Liberal Arts and Sciences (in particular, J. Ward, T. Gabay, and composition and microhabitats of frogs within the Arakan T. Falicov), which supported attendance of Brown lab members at Valley Conservation Area, Cotobato, Mindanao Island, the 2019 International Biogeography Society meetings (Malaga, Philippines. Banwa 3: 16–30. Spain), thus facilitating visits by RMB and JGV to MNCN. Thanks Ferner, J.W., Brown, R.M., Sison, R.V., Kennedy, R.S. (2000): The to P.L. Wood, Jr., for assistance with graphics in Fig. 1. We thank amphibians and reptiles of Island, Philippines. Asiatic H. Kaiser for providing detailed advice regarding nomenclatural Herpetological Research 9: 34–70. technicalities, J. Bernstein for his attentive editing efforts, G. Gaulke, M. (2011): The Herpetofauna of Panay Island, Philippines. Guerra, and S. Mecke for constructive reviews of earlier drafts Frankfurt am Main, Germany, Edition Chimaira. of this manuscript. Gogorza, J. (1888): Datos para la fauna filipina. Vertebrados. Anales de la Sociedad Española de Historia Natural 17: 247–303. References González-Fernández, J.E. (2002) Colección de anfibios y reptiles. In: Catálogo de las colecciones zoológicas de Asia del Museo Alcala, A.C. (2004a): A life of service to science: a tribute to Walter Nacional de Ciencias Naturales, III: Vertebrados. Manuales Creighton Brown 1913–2002. Silliman Journal 45: 258–262. Técnicos de Museología, Museo Nacional de Ciencias Alcala, A.C. (2004b): Biodiversity research in the Philippines Naturales, Madrid 13: 61–180. from 1997–2003. ASEAN Biodiversity 4: 26–31. Günther, A. (1859): On the genus Elaps of Wagler. Proceedings of Bogert, C.M. (1943): Dentitional phenomena in cobras and other the Zoological Society of London 27: 79–93. elapids with notes on adaptive modifications of fangs. Bulletin Günther, A. (1864): The Reptiles of British . London, United of the American Museum of Natural History 81: 285–360. Kingdom, Taylor & Francis. xxvii + 452 pp. Brown, R.M., Diesmos, A.C. (2009): Philippines, Biology. In: Leviton, A.E. (1963): Remarks on the zoogeography of Philippine Encyclopedia of Islands, p.723–732, Gillespie, R., Clague, terrestrial snakes. Proceedings of the California Academy of D., Eds., Berkeley, California, United States, University of Sciences 42: 112–145. California Press. Leviton, A.E. (1964): Contributions to a review of Philippine Brown, R.M., McGuire, J.A., Ferner, J.W., Icarangal, N., Jr., snakes. III. The genera Maticora and Calliophis. Philippine Kennedy, R.S. (2000): Amphibians and reptiles of Luzon Journal of Science 92: 523–550. First Report of Calliophis salitan from Mindanao Island 1035

Leviton, A.E., Brown, R.M., Siler, C.D. (2014): The dangerously Wallach, V., Brown, R.M., Diesmos, A.C., Gee, G.V.A. (2007): venomous snakes of the Philippine Archipelago. In: The Coral An enigmatic new species of blind snake from Luzon Island, Triangle: the 2011 Hearst Biodiversity Philippine Expedition, p. northern Philippines (Serpentes: Typhlopidae). Journal of 473–530, Williams, G.C., Gosliner, T.M, Eds., San Francisco, Herpetology 41: 690–702. California, USA, California Academy of Sciences. Watt, G., Padre, L., Tuazon, M.L., Theakstron, R.D.Z., Laughlin, L. Leviton, A.E, Siler, C.D., Weinell, J.L., Brown, R.M. (2018): A (1988): Bites by the Philippine Cobra Naja naja philippinensis: synopsis of the snakes of the Philippines: a synthesis of data prominent neurotoxicity with minimal local signs. American from biodiversity repositories, field studies, and the literature. Journal of Tropical Medicine and Hygiene 39: 306–311. Proceedings of the California Academy of Sciences 64: 399–568. Weinell, J.L., Hooper, E., Leviton, A.E., Brown, R.M. (2019): Nuñeza, O., Ates, F.B., Alicante, A.A. (2010): Distribution of Illustrated key to the snakes of the Philippines. Proceedings of endemic and threatened herpetofauna in Mt. Malindang, the California Academy of Sciences 66: 1–49. Mindanao, Philippines. Biodiversity Conservation 19: 503–518. Weinell, J.L., Paluh, D.J., Siler, C.D., Brown, R.M. (2020): A new, Plettenberg Laing, A. von (2018): A multilocus phylogeny of miniaturized genus and species of snake (Cyclocoridae) from the cobra clade elapids. Unpublished PhD thesis, Bangor the Philippines. Copeia 108: 907–923. University, Bangor, Wales, United Kingdom. Wüster, W., Thorpe, R.S. (1991): Asiatic cobras: systematics and Rabor, D.S., Alcala, A.C. (1959): Notes on a collection of amphibians snakebite. Experientia 47: 205–209. from Mindanao Island, Philippines. Silliman Journal 2: 93–102. Wuster, W., Thorpe, R.S. (1992): Dentitional phenomena in cobras Ross, C.A., Lazell, J. (1991): Amphibians and reptiles of Dinagat revisited: fang structure and spitting in the Asiatic species of and Siargao islands, Philippines. Philippine Journal of Science Naja (Serpentes: Elapidae). Herpetologica 48: 424–434. 119: 257–286. Wynn, A., Diesmos, A.C., Brown, R.M. (2016): Two new Sanguila, M.B., Cobb, K.A., Siler, C.D., Diesmos, A.C., Alcala, species of Malayotyphlops from the northern Philippines, with A.C., Brown, R.M. (2016): The amphibians and reptiles of redescriptions of M. luzonensis (Taylor) and M. ruber (Boettger). Mindanao Island, southern Philippines, II: the herpetofauna of Journal of Herpetology 50: 157–168. northeast Mindanao and adjacent islands. Zookeys 624: 1–132. Siler, C.D., Welton, L.J., Siler, J.M., Brown, J., Bucol, A., Diesmos, A.C., Brown, R.M. (2011): Amphibians and reptiles, Luzon Island, Aurora Province and Aurora Memorial National Park, northern Philippines: new island distribution records. Check List 7: 182–195. Sison, R., Gonzales, P.C., Ferner, J.W. (1995): New records from Panay, Philippines. Herpetological Review 26: 48–49. Slowinski, J.B., Boundy, J., Lawson, R. (2001): The phylogenetic relationships of Asian coral snakes (Elapidae: Calliophis and Maticora) based on morphological and molecular characters. Herpetologica 57: 233–245. Smith, B.E. (1993a): Notes on a collection of squamate reptiles from eastern Mindanao, Philippines Islands, Part 1: Lacertilia. Asiatic Herpetological Research 5: 85–95. Smith, B.E. (1993b): Notes on a collection of squamate reptiles from eastern Mindanao, Philippines Islands, Part 2: Serpentes. Asiatic Herpetological Research 5: 95–102. Supsup, C.E., Guinto, F.M., Redoblado, B.R., Gomez, R.S. (2017): Amphibians and reptiles from the Mt. Hamiguitan Range of eastern Mindanao Island, Philippines: new distribution records. Check List 13: 1–14. Stuebing, R.B, Inger, R.F., Larder, B. (2014): A Field Guide to the Snakes of Borneo. Kota Kinabalu, Malaysia, Natural History Publications (Borneo). Taylor, E.H. (1922): The Snakes of the Philippine Islands. Manila, Philippines, Bureau of Printing. Taylor, E.H. (1975): Philippine adventures: an autobiographical memoir. In: Recollections of an Herpetologist, p. 1–105, Taylor, E.H., Leonard, A.B., Smith, H.M., Pisani, G.R., Eds., Lawrence, Kansas, United States, University of Kansas Museum of Natural History (Monograph No. 4). Uetz, P., Freed, P., Hošek, J. (2020): The Database. Accepted by Justin Bernstein Available at http://www.reptile-database.org. Accessed on 29 December 2020.